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Species co-occurrence, nestedness and guild–environment relationships in stream macroinvertebrates



    1. Finnish Environment Institute, Research Programme for Biodiversity, University of Oulu, Oulu, Finland
    2. Department of Biology, University of Oulu, Oulu, Finland
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Jani Heino, Finnish Environment Institute, Research Programme for Biodiversity, PO Box 413, University of Oulu, FI-90014 Oulu, Finland. E-mail:


1. Describing species distribution patterns and the underlying mechanisms is at the heart of ecological research. A number of recent studies have used null model approaches to explore mechanisms behind spatial variation in community structure.

2. However, unexplored questions are the degree to which single guilds of potentially competing stream macroinvertebrate species show: (i) interspecific segregation among-stream sites (i.e. occur together less often than expected by chance), suggesting competitive interactions; (ii) interspecific aggregation (i.e. occur together more often than expected by chance), suggesting similar responses to the environment; (iii) comply with nestedness, suggesting the existence of selective extinctions or colonisations and (iv) show similar environmental relationships.

3. The present analyses showed that guilds of stream macroinvertebrates exhibit non-random co-occurrence patterns that were generally contingent on the weighting of sites by stream size. Despite significant segregation of species, each guild also showed significantly nested patterns. Species richness was correlated with different environmental factors between the guilds, although these correlations were relatively low. By contrast, correlations between the major ordination axes and key environmental variables were slightly stronger in canonical correspondence analysis, and generally the same factors were most strongly correlated with variation in the species composition of each guild.

4. The present findings are the first to show that species within each stream macroinvertebrate guild show significant negative co-occurrence at the among-stream riffle scale. These findings present challenges for future studies that aim to disentangle whether these patterns comply with the habitat checkerboard or the competitive checkerboard explanations.

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